1Descriptions of new Australian chondrichthyans

Description of two new species of gulper sharks, genus Centrophorus (Chondrichthyes: Squaliformes: Centrophoridae) from Australia

William T. White1, David A. Ebert2 & Leonard J.V. Compagno3

1CSIRO Marine & Atmospheric Research, GPO Box 1538, Hobart, TAS, 7001, AUSTRALIA2PacificSharkResearchCenter,MossLandingMarineLaboratories,MossLanding,CA95039,USA

3Shark Research Center, Iziko – Museums of Cape Town, Cape Town 8000, SOUTH AFRICA

ABSTRACT.— Two new species of gulper sharks, Centrophorus westraliensis sp. nov. and Centrophorus zeehaani sp. nov., are described based on specimens from Australian waters. Centrophorus westraliensis waspreviouslyconsideredtobeconspecificwithC. harrissoni from eastern Australia, New Zealand and New Caledonia. However, recent investigations revealed that the two forms are clearly separable based on their morphology. Centrophorus zeehaaniwaspreviouslyconsideredtobeconspecificwithCentrophorus uyato, but the original description of the latter species clearly depicts a Squalus and not a Centrophorus species. Centrophorus zeehaani belongs to a subgroup of longnose Centrophorus species which includes C. harrissoni, C. isodon, C. seychellorum and C. tessellatus, and is clearly separable based on morphology and coloration. The two new species appear to be endemic to Australian waters and their conservation status needs to be investigated to determine if they are critically endangered as is the case with C. harrissoni.

Key words. Centrophorus westraliensis – Centrophorus zeehaani – Centrophorus harrissoni – new species – endemic – Australia

PDF contact: john.pogonoski@csiro.au

INTRODUCTION

The Centrophoridae (Chondrichthyes: Squaliformes) comprises two genera, Centrophorus and Deania, which consist of ten and four species, respectively (Compagno et al., 2005). The genus Centrophorus, proposed by Müller & Henle, 1837, has had a very complex taxonomic history. Thisisdue,inpart,tothesubstantialintraspecificvariationin body and fin morphology and ontogenetic changesin dermal denticle and tooth morphologies associated with growth and sexual dimorphism. Compounding the identificationproblem is thatmost specieswerepoorlydescribed originally and many of the characteristics usedtodistinguishbetweenspecieswerepoorlydefined.Furthermore, the poor representation of Centrophorus species in regional institutional collections, due in part to their large size and occurrence in deep benthic habitats, has also been a hindrance.

Last & Stevens (1994) provided treatments for fivespecies of Centrophorus occurring in Australian waters, i.e. Centrophorus granulosus (Bloch & Schneider, 1801), C. harrissoni McCulloch, 1915, C. moluccensis Bleeker, 1860, C. squamosus (Bonnaterre, 1788) and C. uyato(Rafinesque,1810).Theauthorsnotedthateastand west coast populations of C. harrissoni required closer comparative examination. Compagno et al. (2005) listedthefirstfouraforementionedspeciesasoccurring

in Australian waters, plus possibly C. atromarginatus Garman, 1913 and C. niaukang Teng, 1959, but commented that C. uyato was not a valid taxon. Other Centrophorus species recognised by Compagno et al. (2005) which occur outside of Australia are C. acus Garman, 1906, C. isodon (Chu, Meng & Liu, 1981), C. lusitanicus Bocage & Capello, 1864 and C. tessellatus Garman, 1906. Although many authors have included Rafinesque’s (1810) Squalus uyatus in the genus Centrophorus following Garman (1906), the original description and illustration (Rafinesque, 1810: 13–14,pl. 14, fig. 2) clearly depicts a Squalus species that is recognisable by its sharp snout, broad, slightly attenuated, narrowlyroundedpectoralfreereartip,largefirstdorsalfin spine, second dorsal fin much lower than the firstand with a deeply incised posterior margin, very large claspers with prominent spines or spurs (very small in Centrophorus and with relatively inconspicuous spurs), a short abdomen, an elongated precaudal tail and long caudal peduncle, a deeply notched postventral caudal margin, and no subterminal notch. However, despite the invalid species name, the Australian Centrophorus species treated as C. uyato by Last & Stevens (1994) is certainly a valid taxon. The validity of the most recent described Centrophorus species, C. seychellorum Baranes, 2003, needs to be resolved.

The critical importance of gaining a better understanding

2

of the taxonomic status and distributions of Centrophorus species occurring in Australian waters is highlighted by the severe population declines reported off southeastern Australiaasaresultofanintensivetrawlfishery.Grahamet al. (2001) reported declines of 98.4–99.7% in the relative abundances of C. harrissoni, C. moluccensis and C. cf. uyato off the upper slope of New South Wales between 1976–77 and 1996–97. As a result of these serious declines, these three species have been listed as either endangered or critically endangered by the IUCN Red List of Threatened Animals (www.redlist.org). Therefore, given the severely threatened status of some Centrophorus species and populations in Australian waters it is critical that taxonomic issues for this genus are resolved so that accurate identification and propermanagement policies for these sharks can be implemented. Here we describe two new species of Centrophorus, both of which appear to be Australian endemics, redefineCentrophorus harrissoni, and compare these species with other closely related species of this genus. Collections of Centrophorus species from the Western Indian Ocean, fromeasternIndonesianfish landingsitesandmarkets,and from Taiwan and the Philippines has allowed for comparison with the new species, and other Australian representatives of this genus. The nomenclature of Australian Centrophorus is also discussed.

METHODS

The morphometric measurements used generally followed those detailed by Compagno (1984, 2001), but focused on direct (point-to-point) rather than horizontal measurements. Data in the literature is often unreliable for comparative purposes due to the different and unspecifiedmethodology adopted.Measurements weretaken following the methodology described by Last et al. (2007) for the genus Squalus with some additional measurements, i.e. CST — subterminal caudal-finmargin, CTL — terminal caudal-fin lobe, DPI — 1st dorsal-finmidpointtopectoral-fininsertion,D1SL—1st dorsalsoftfinlength(fromperpendiculartojunctionofexposed spineand softfinbase to free rear tip),D2SL— 2nd dorsal soft fin length (from perpendicular tojunctionofexposedspineandsoftfinbase to free reartip), DPO — 1stdorsal-finmidpointtopelvic-finorigin,PDI—pelvic-finmidpointto1stdorsalfininsertion,andPDO—pelvic-finmidpointto2nddorsal-finorigin,fromCompagno (2001).

All type specimens of the two new species were measured in full and compared with 7 measured specimens of C. harrissoni (CSIRO CA 4103, CSIRO H 866–01, CSIRO H 866–05, CSIRO H 866–06, CSIRO H 2528–01, CSIRO T 810 and CSIRO H 6310–05) and 5 specimens of C. isodon (CSIRO H 5857–01, CSIRO H 5889–15, CSIRO H 5875–04, CSIRO H 6125–02, and CSIRO H 6138–01) (Tables 1–3). A subsample of measurements were also taken from the holotype of

C. tessellatus (MCZ 1031S). Vertebral counts were obtained from radiographs of all of the type specimens of the two new species, and comparative vertebral meristics were obtained from radiographs or from dissections of fresh material of C. harrissoni and C. isodon. Counts were obtained separately for trunk (monospondylous precaudal centra), precaudal (monospondylous precaudal centra + diplospondylous precaudal centra to origin of the caudal-fin upper lobe) and diplospondylous caudalcentra (centra of the caudal fin) vertebrae. Tooth rowcounts were taken in situ,bymakingincisionsatthejawangles to expose the teeth, or from preserved jaws. Inthe descriptions of the new species, morphometric and meristicvaluesfortheholotypearegivenfirst,followedin parentheses by the ranges of the paratypes.

Material examined, including type specimens of the new species, are deposited in the Australian National Fish Collection, Hobart, Australia (CSIRO), and ichthyological collections of the Australian Museum, Sydney (AMS), Museum Victoria, Melbourne (NMV), Museum of Comparative Zoology, Harvard (MCZ) and the Queensland Museum, Brisbane (QM).

Centrophorus harrissoni McCulloch, 1915

Figs 1, 2a, 3a, Table 1

Material examined. 19 whole specimens: AMS E–5570 (Holotype) eviscerated female 736 mm TL, Gabo Island, Victoria, Australia; CSIRO CA 4103, adult male 843mmTL,northofFlindersIsland,BassStrait,39°05′S,148°38′E,444–468m,06May1984;CSIROH866–01, female 420 mm TL, CSIRO H 866–05, immature male 350 mm TL, CSIRO H 866–06, immature male 417mmTL,eastofJervisBay,NewSouthWales,34°58′S,151°09′E,490–576m,10Sep1986;CSIROH987–01, immature female ca. 390 mm TL, north of Flinders Island,BassStrait,39°02′S,148°37′E,440–448m,29Nov 1984; CSIRO H 2528–01, female 1049 mm TL, east ofMariaIsland,Tasmania,42°39′S,148°28′E,500m,15 July 1990; CSIRO H 6310–05, adult male 874 mm TL, north-east of Flinders Island, Tasmania, 39°00′ S,148°38′E,500–680m;CSIROH6498–03,adultmale825 mm TL, CSIRO H 6498–04, adult male 925 mm TL, southern half of Flinders Island, Tasmania, 300–500 m, 24 June 2003; CSIRO H 6500–03, adult male 910 mm TL, offFlindersIsland,Tasmania,40°15′S,148°45′E,329– 512 m, 21 Aug 2003; CSIRO H 6501–01, female 1114 mm TL, CSIRO H 6501–02, adult male 882 mm TL, CSIRO H 6501–03, adult male 907 mm TL, east of Flinders Island, Tasmania, ca. 41° S, ca. 149° E, 403–439 m, 18 June 2004; CSIRO H 6502–01, adult male 890 mm TL, CSIRO H 6502–02, female 940 mm TL, south of South EastCape,Tasmania,43°45′S,146°45′E,329–549m,27Aug 2002; CSIRO T 810, female 850 mm TL, Oct 1981, off St Helens, Tasmania, 580 m; QM I 35759, immature male 387 mm TL, QM I 35770, female 600 mm TL, Fraser

3Descriptions of new Australian chondrichthyans

Figure 1. Lateral view of Centrophorus harrissoni: A. CSIRO H 2528–01 (female 1049 mm TL); B. CSIRO H 987–01 (immature female ca. 390 mm TL).

Seamount, Queensland, 24°25′ S, 155°17′ E, 670 m. 18 skeletal specimens: CSIRO H 6307–03, immature male 557 mm TL, CSIRO H 6307–04, female 1025 mm TL, CSIRO H 6307–05, immature male 565 mm TL, CSIRO H 6307–06, female 933 mm TL, CSIRO H 6307–07, adult male 891 mm TL, east of Flinders Island, Tasmania, ca. 40° S, ca. 149° E, 350–430 m, 12 July 2004; CSIRO H 6308–01, adult male 900 mm TL, CSIRO H 6308–02, female 1039 mm TL, CSIRO H 6308–03, female 716 mm TL, Banks Strait, Tasmania, ca. 40° S, ca. 148° E, 29 July 2004; CSIRO H 6309–03, adult male 902 mm TL, CSIRO H 6309–05, adult male 939 mm TL, east of Flinders Island, Tasmania, ca. 40° S, ca. 149° E, 400– 450 m, 01 Aug 2004; CSIRO H 6310–01, adult male 926 mm TL, CSIRO H 6310–03, female 870 mm TL, north-eastofFlindersIsland,Tasmania,39°00′S,148°38′E, 500–680 m, 24 July 1986; CSIRO H 6498–03, male 825 mm TL, southern half of Flinders Island, Tasmania, 300–500 m, 24 June 2003; CSIRO H 6499–01, female 1080 mm TL, CSIRO H 6499–02, female 880 mm TL, CSIRO H 6499–03, female 1070 mm TL, CSIRO H 6499–04, female 778 mm TL, north-east coast of Tasmania, ca. 41° S, 149° E, 24 July 2003; CSIRO H 6500–01, adult male902mmTL,offFlindersIsland,Tasmania,40°15′S,148°45′E,329–512m,21Aug2003.

DIAGNOSIS.— A moderate-sized species of Centrophorus with the following combination of adult characters: pre-second dorsal length 61.9–63.2% TL, 6.6–8.4timesdorsal–caudalspace;pre-firstdorsallength30.4–32.0% TL; interdorsal space 18.2–20.6% TL, 2.2–2.6 times dorsal–caudal space; dorsal–caudal space 7.5–

9.4% TL, 3.2–4.0 in pectoral–pelvic space; head long and robust (length 22.4–24.6% TL, 2.7–3.0 times mouth width; width 13.2–14.0% TL, 4.5–4.8 in pre-second dorsal length; width at anterior of nostrils 7.7–7.9% TL); snout long (preoral length 11.4–12.4% TL, 2.0–2.2 times head height at anterior of mouth, 1.3–1.5 times mouth width; horizontal preorbital length 7.1–8.2% TL; horizontal prenarial length 4.4–5.4% TL); mouth large (width 7.8–8.5%TL);pectoralfinmoderatelylarge(anteriormargin11.9–12.4%TL, 2.3–2.4 times base length); caudal finlarge (dorsal caudal margin 19.1–19.5% TL; 2.1–2.5 times dorsal–caudal space); first dorsal fin moderatelylarge and tall (height 6.2–7.0% TL), spine moderately robust(basewidth0.9–1.0%TL).Dorsalfinsofjuvenileswith a distinct blackish oblique blotch anteriorly and a white blotch on the upper posterior margin; adults with a less distinct dark blotch (usually still apparent in fresh specimens) and with a white blotch restricted to a narrow white posterior margin (occasionally indistinct in largest specimens). Flank denticles of adults flat,block-like, not overlapping, crenulate. Upper teeth of females and immature males strongly oblique, similar in shape, but much smaller than, lower teeth; upper teeth of mature males erect, upright, becoming only slightly oblique laterally. Tooth row count: 37–39/30 or 31 (n=3). Total vertebral centra 117–126 (mean 121.6, n=20), monospondylous precaudal centra 53–59 (56.4, n=21), diplospondylous precaudal centra 29–37 (32.7, n=21) and precaudal centra 85–94 (89.1, n=21).

SIZE.— Specimens examined ranged from 350 to 1114 mm TL. Smallest mature male was 825 mm TL.

A

B

4

Table 1. Proportional dimensions as percentages of total length for the holotype (AMS E 5570; taken from Duffy, 2007), 4adultsand3juvenilesofCentrophorus harrissoni,and3adultsand2juvenilesofCentrophorus isodon.

C. harrissoni C. isodon

Holotype Adults Juveniles Adults JuvenilesMin. Max. Min. Max. Min. Max. Min. Max.

Total length (mm) 736 843 1049 350 420 952 989 530 540Precaudal length 78.4 80.2 80.6 76.7 77.6 81.5 82.6 78.9 79.6

Pre-second dorsal length 61.0 61.9 63.2 59.4 61.2 66.2 68.0 63.0 64.3Pre-firstdorsallength 32.3 30.4 32.0 31.8 33.0 30.8 31.9 30.2 31.8Pre-vent length 56.2 58.4 60.1 55.4 55.6 60.1 61.1 57.0 58.9Prepelvic length 54.6 56.2 57.4 52.6 53.8 58.3 59.7 54.9 57.4Prepectoral length 21.7 22.0 23.8 24.6 26.0 20.9 22.8 24.1 24.2Head length 23.5 22.4 24.6 25.5 26.2 22.0 23.3 24.2 24.8Prebranchial length 20.2 19.7 20.5 22.3 22.6 18.3 19.8 21.1 21.3Prespiracular length 14.5 14.1 14.9 15.9 16.2 12.3 13.2 14.4 14.6Preorbital length 7.5 7.5 8.6 9.4 9.5 6.7 7.7 8.1 8.3Prenarial length 4.9 4.9 5.8 5.8 6.2 4.4 4.9 4.8 5.5Preoral length 13.2 11.4 12.4 13.0 13.5 10.1 10.8 11.0 11.9Inner nostril–labial furrow space 7.1 6.2 6.4 7.2 7.8 5.9 6.3 6.0 6.5Mouth width 7.3 7.8 8.5 7.9 8.3 7.4 7.9 8.1 8.5Upper labial furrow length 2.2 2.3 2.7 2.0 2.5 1.8 2.1 1.9 2.3Nostril width 1.9 1.7 1.9 2.0 2.3 1.6 1.9 2.0 2.0Internarial space 4.6 3.8 4.4 4.3 4.7 3.3 3.9 3.5 4.3Interorbital space 8.7 7.3 8.1 7.4 8.5 7.2 7.5 7.5 7.9Eye length 5.7 5.7 6.6 6.9 7.7 5.7 5.9 5.9 6.5Eye height 1.5 1.6 2.0 2.1 2.8 1.7 1.9 1.4 2.0Spiracle diameter - greatest 1.5 1.3 1.7 1.4 2.1 1.7 1.9 2.0 2.1First gill-slit height 2.4 1.6 2.5 1.9 2.4 1.7 1.9 2.1 2.1Fifth gill-slit height 1.8 2.2 2.4 2.1 2.8 2.5 2.7 2.5 2.7Interdorsal space 18.4 18.2 20.6 16.1 19.7 22.4 25.1 21.1 21.1Dorsal–caudal space 8.6 7.5 9.4 8.4 8.9 5.8 6.6 6.1 7.4Pectoral–pelvic space 26.9 29.5 31.9 25.0 26.3 32.1 33.3 29.1 29.5Pelvic–caudal space 17.0 13.6 15.3 13.7 15.1 14.3 15.0 14.9 15.6First dorsal length – 18.0 19.6 16.7 18.2 18.8 19.8 18.6 19.2Firstdorsalsoftfinlength 12.0 11.4 11.9 11.0 11.5 12.0 12.6 12.2 13.0First dorsal anterior margin 11.1 11.5 13.5 11.5 13.4 12.2 13.5 12.6 12.6First dorsal base length 11.0 12.7 14.1 10.6 12.7 13.8 14.3 12.3 13.2First dorsal height 7.8 6.2 7.0 5.2 6.2 5.7 6.2 5.7 6.8First dorsal inner margin 6.4 6.0 6.2 5.6 6.3 5.5 6.0 5.9 6.8First dorsal posterior margin 10.4 9.2 10.9 7.8 10.4 9.1 9.8 9.4 10.3First dorsal spine length 1.0* 2.4 2.6 1.7 1.8 1.8 2.3 2.1 2.7First dorsal spine base width – 0.9 1.0 0.8 0.9 0.7 0.7 0.6 0.8Second dorsal length – 13.7 14.7 12.7 14.1 13.4 13.9 14.2 14.3Seconddorsalsoftfinlength 9.5 8.3 8.8 8.1 8.7 7.8 8.6 8.7 9.0Second dorsal anterior margin 9.1 8.5 10.0 8.6 10.1 9.1 9.6 9.5 9.8Second dorsal base length 9.2 9.2 10.1 8.0 9.7 9.4 9.7 9.1 9.8Second dorsal height 5.0 4.8 5.2 3.8 4.5 4.4 4.8 4.6 5.4Second dorsal inner margin 4.9 4.1 4.7 4.7 5.3 4.1 4.4 4.3 5.3

* denotes possible damage resulting in underestimate of spine length

5Descriptions of new Australian chondrichthyans

C. harrissoni C. isodon

Holotype Adults Juveniles Adults JuvenilesMin. Max. Min. Max. Min. Max. Min. Max.

Second dorsal posterior margin 8.1 7.4 8.2 6.8 7.4 6.6 7.2 7.0 7.4Second dorsal spine length 2.1 1.6 2.6 2.2 2.3 1.7 2.2 2.7 3.1Second dorsal spine base width – 0.7 0.8 0.6 0.9 0.6 0.7 0.8 0.8Pectoral anterior margin 13.0 11.9 12.6 11.4 12.8 11.1 12.2 13.5 13.5Pectoral inner margin 10.0 11.6 14.1 10.6 11.4 11.5 12.6 11.8 11.8Pectoral base length – 5.1 5.2 4.0 4.4 4.5 4.7 5.0 5.0Pelvic length 10.0 9.9 11.2 9.6 10.1 7.6 10.4 9.8 10.3Pelvic height 5.8 5.8 6.5 4.1 4.9 5.2 6.1 4.3 5.6Pelvic inner margin 5.4 5.0 6.3 4.9 5.3 5.6 6.4 5.1 5.5Dorsal caudal margin 21.3 19.1 19.5 21.2 23.1 16.4 18.0 20.9 21.1Preventral caudal margin 11.7 11.6 14.1 13.2 15.4 10.8 12.3 12.8 13.1Upper postventral caudal margin – 6.8 7.9 8.4 9.2 7.0 8.3 6.6 7.9Lower postventral caudal margin – 3.1 4.7 3.3 3.6 4.0 4.9 3.1 4.0

Caudal fork width – 6.1 7.2 6.9 7.4 6.2 7.0 6.3 6.9Caudal fork length – 11.3 13.0 13.2 16.0 10.4 11.7 12.8 13.3Caudal terminal lobe 9.1 8.2 8.7 7.4 8.2 6.5 7.8 7.3 7.4Caudalsubterminalfinmargin – 2.1 2.6 2.9 3.7 1.5 2.6 3.3 3.7Head width at anterior of nostrils 7.6 7.7 7.9 8.6 9.2 6.8 7.2 8.2 8.3Head width at mouth 11.2 10.2 10.7 10.6 11.3 8.8 9.2 8.9 10.9Head width 13.4 13.2 14.0 10.0 13.1 9.7 11.9 9.2 11.2Trunk width – 9.8 11.3 6.9 9.6 – – – –Abdomen width – 10.5 11.7 6.1 8.1 – – – –Tail width – 5.1 6.6 3.8 4.8 – – – –Caudal peduncle width – 2.7 3.3 2.3 2.8 3.0 3.2 2.2 2.3Head height at mouth – 5.6 5.7 5.9 6.8 – – – –Head height – 9.8 11.0 8.6 10.2 8.1 9.5 7.1 8.7Trunk height – 11.5 12.6 8.5 11.0 – – – –Abdomen height – 11.8 12.5 8.1 10.9 – – – –Tail height – 6.7 7.1 6.6 7.2 – – – –Caudal peduncle height – 3.4 3.8 3.5 3.8 3.0 3.2 3.3 3.5Clasper outer length – 2.9 3.5 0.7 1.0 – – – –Clasper inner length – 6.8 7.7 2.7 2.7 – – – –Clasper base width – 1.0 1.0 0.6 0.7 – – – –First dorsal midpoint–pectoral insertion – 8.4 9.7 8.9 10.4 10.0 11.0 8.3 9.5First dorsal midpoint–pelvic origin – 17.9 20.6 13.4 15.4 21.9 23.3 18.8 19.5Pelvicmidpoint–firstdorsalinsertion – 13.8 16.5 11.0 13.3 16.0 18.6 14.6 14.8Pelvic midpoint–second dorsal origin – 3.6 5.2 4.7 6.1 4.9 6.2 5.3 5.6

Table 1. cont’d.

DISTRIBUTION.— Known from the upper to mid-continental slope of eastern Australia from Clarence River (ca.29°S)toSouthEastCape,Tasmania(43°45′S),andfrom Fraser Seamount,Queensland (24°25′ S, 155°17′E) in depths of 300–680 m (Fig. 11). It has also been recorded from theKermadecRidge (31°45′S, 178°52′

E),NorfolkRidge(33°07′S,166°52′E)andThreeKingsRidge(31°15′S,172°56′E)offNewZealandindepthsof 536–710 m (Duffy, 2007) and possibly New Caledonia (Compagno et al., 2005; Anon., 1998). Last & Stevens (1994) recorded this species at depths of 220–790 m.

6

Figure 2. Ventral view of the head of: A. Centrophorus harrissoni (CSIRO H 2528–01, female 1049 mm TL); B. Centrophorus isodon (not retained, female 1010 mm TL,Kedongananfishmarket,Bali,Indonesia).

A

B

Figure 3. Cusps of the flank denticles of: A. Centrophorus harrissoni (CSIRO H 6309–03, adult male 980 mm TL); B. Centrophorus isodon (CSIRO H 6125–02, female 965 mm TL).

Longnose Centrophorus similar in general morphology and coloration to C. harrissoni are known from off Taiwan, the east coast of South Africa, and the Western North Atlantic but their status needs detailed investigation (Compagno et al., 2005).

VERNACULAR.—Harrisson’sDogfish.

REMARKS.— Centrophorus harrissoni is most similar in appearance to C. isodon (Figs 2b, 3b, 4) and C. tessellatus, which together form a subgroup of longnose Centrophorus species. Adults of C. harrissoni differ from those of C. isodoninhavingmuchcloserdorsalfins(interdorsal space 18.2–20.6 vs. 22.4–25.1% TL, 2.2–2.6 vs. 3.4–4.3 times dorsal–caudal space); a longer caudal fin (precaudal length 80.2–80.6 vs. 81.5–82.6% TL,dorsal caudal margin 19.1–19.5 vs. 16.4–18.0% TL); a larger mouth (width 7.8–8.5 vs.7.4–7.9% TL); a broader head (head width 13.2–14.0 vs. 9.7–11.9% TL; 4.5–4.8 vs. 5.7–6.9 in pre-second dorsal length); the upper teeth of females far more oblique; flank denticles with lowbut obvious ridges (vs. no obvious ridges; Fig. 3b) and

dorsal fins with a whitish posterior margin in all sizeclasses (except some very large specimens). Juveniles of C. harrissoni differ from those of C. isodon in the following measurements and ratios: pre-second dorsal length 59.4–61.2 vs. 63.0–64.3% TL; pre-vent length 55.4–55.6 vs. 57.0–58.9% TL; dorsal–caudal space 8.4–8.9 vs. 6.1–7.4% TL; pectoral–pelvic space 25.0–26.3 vs. 29.1–29.5% TL, 2.9–3.0 vs. 3.9–4.8 times dorsal–caudal space; preoral length 13.0–13.5 vs. 11.0–11.9% TL; inner nostril to upper labial furrow 7.2–7.8 vs. 6.0–6.5% TL; firstdorsal-finsoftlength11.0–11.5vs.12.2–13.0%TL;midfirstdorsal-finbasetopelvic-finoriginspace13.4–15.4 vs. 18.8–19.5% TL.

Centrophorus harrissoni differs from C. tessellatus in having a longer pectoral–pelvic space (29.5–31.9 vs. 27.9% TL); shorter pelvic–caudal space (13.6–15.3 vs. 17.9% TL); a slightly larger mouth (7.8–8.5 vs. 7.4% TL); a broader head (head width 13.2–14.0 vs. 12.2% TL) and no white borders on the gill slits.

A

B

7Descriptions of new Australian chondrichthyans

Figure 4. Lateral view of Centrophorus isodon (not retained, female1010mmTL,Kedongananfishmarket,Bali,Indonesia).

The upper teeth of Centrophorus harrissoni show strong sexual dimorphismwith those of females and juvenilemales being strongly oblique (n=8) whilst those of mature males being erect and upright, and only slightly oblique laterally (n=5). The lower teeth of C. harrissoni also display sexual dimorphism, although generally not as marked as that in the upper teeth, with those of the adult males being strongly oblique and blade-like, as in females, but with the tip curved upwards. This sexual dimorphism was also recorded and illustrated for this species by Duffy (2007) in New Zealand waters. This difference in upper tooth shape was also noted by Baranes (2003) for C. seychellorum, with those of the female holotype possessing strongly oblique cusps and those of the adult male paratype possessing upright cusps (Fig. 22 in Baranes, 2003). The validity of this species has not yet been investigated and this issue is not helped bythefact that theholotypehasadamagedcaudalfin.Since all morphometric measurements are expressed as percentage of total length, they are not accurate for the holotype and thus not easily comparable to other species and so comparisons made below are based on the paratype. The preoral length of this species (10.1% TL) is smaller than in C. harrissoni (11.4–12.4% TL), but similar to C. isodon (10.1–10.8% TL) and C. tessellatus (10.1% TL). The upper teeth of the female holotype of C. seychellorum are far more oblique than in females of C. isodon, and these two species also differ in dorsal–caudal space (8.2 vs. 5.8–6.6% TL). Centrophorus seychellorum differs from C. tessellatus in having a shorter head (21.5 vs. 24.8% TL); larger interdorsal space (22.9 vs. 19.4% TL) and a shorter horizontal prenarial length (2.2vs.3.7%TL).TheflankdenticlesofC. seychellorum arealsomorepointedandclosertogetherthantheflattermore block-like denticles of C. isodon, C. tessellatus and C. harrissoni (see Figs 23 and 24 in Baranes, 2003). Thus, C. seychellorum is potentially a valid species belonging to the longnose subgroup, but further investigation of this species is required.

Centrophorus westraliensis sp. nov.

Figs 5–7, Table 2

Centrophorus harrissoni (in part): Last & Stevens, 1994: pp 52, 56, 57; Daley, Stevens, Last & Yearsley, 2002: pp 73.

Holotype. CSIRO H 2625–06, female 909 mm TL, west ofPointD’Entrecausteaux,WesternAustralia,35°00′S,114°42′E,738–750m,18Feb1991.Paratypes. 5 whole specimens: CSIRO H 2357–03, female 789 mm TL, south of Cape Leeuwin, Western Australia, 35°02′ S, 115°02′ E, 673 m, 23 Dec 1989;CSIRO H 2358–01, immature male 371 mm TL, north-westofGeraldton,WesternAustralia,28°13′S,113°07′E, 616 m, 27 Dec 1989; CSIRO H 2580–01, female 414 mm TL, south-west of Shark Bay, Western Australia, 27°05′S,112°22′E,713–714m,31Jan1991;CSIROH2606–01, female 774 mm TL, west of Rottnest Island, Western Australia, 32°00′ S, 114°55′ E, 640–670 m,10 Feb 1991; CSIRO H 2625–05, female 866 mm TL, collected with holotype.

DIAGNOSIS.— A moderate-sized species of Centrophorus with the following combination of adult characters: pre-second dorsal length 63.8–64.9% TL, 8.8–9.7timesdorsal–caudalspace;pre-firstdorsallength31.3–32.7% TL; interdorsal space 20.3–21.5% TL, 2.8–3.1 times dorsal–caudal space; dorsal–caudal space 6.7–7.3% TL, 4.4–4.9 in pectoral–pelvic space; head long and moderately robust (length 23.2–24.7% TL, 3.1–3.2 times mouth width; width 11.7–12.8% TL, 5.1–5.5 in pre-second dorsal length; width at anterior of nostrils 7.1–7.6% TL); snout long (preoral length 10.9–12.4% TL, 2.0–2.2 times head height at anterior of mouth, 1.4–1.6 times mouth width; horizontal preorbital length 7.0–7.8% TL; horizontal prenarial length 4.5–4.9% TL); mouth moderately large (width 7.6–7.8% TL); pectoral finlarge(anteriormargin12.9–14.2%TL,2.6–2.8timesbase length); caudal fin large (dorsal caudal margin17.4–20.2%TL;2.6–2.9timesdorsal–caudalspace);first

8

Figure 5. Lateral view of Centrophorus westraliensis sp. nov.: A. female holotype (CSIRO H 2625–06, 909 mm TL, preserved); B. female paratype (CSIRO H 2606–01, 774 mm TL, fresh); C. immature male paratype (CSIRO 2358–01, 371 mm TL, preserved).

A

B

C

dorsalfinmoderately-sized(height5.5–6.4%TL),spinerelativelyrobust(basewidth1.1–1.2%TL).Dorsalfinsofjuvenileswithadistinctblackishobliqueblotchanteriorlyand a white blotch on the upper posterior margin; adults with a less distinct dark blotch (usually still apparent in fresh specimens) and with white blotch restricted to a narrow white posterior margin (can be indistinct in larger preserved specimens). Upper teeth of females and of an immature male are strongly oblique, similar in shape, but much smaller than lower teeth. Teeth count: in holotype 38/29.Flankdenticlesofadultsflat,notoverlapping,withscalloped edges. Total vertebral centra 112–117 (mean 115.3), monospondylous precaudal centra 55–57 (55.8), diplospondylous precaudal centra 29–33 (31), precaudal centra 85–88 (86.8) and diplospondylous caudal centra 27–30 (28.5).

DESCRIPTION.— Body fusiform, moderately elongate, nape moderately humped; deepest near firstdorsal-fin spine, trunk height 1.42 (0.98–1.16 in 3adult paratypes) times width, 1.02 (0.76–0.90) times abdomen height; head moderately elongate, length 23.2

(23.8–24.7)% TL; caudal peduncle moderately slender, 16.1 (13.7–14.6)% TL. Head long, moderately robust, broad, width 1.27 (1.23–1.38) times trunk width, 1.17 (1.17–1.45) times abdomen width; depressed forward of spiracles, becoming somewhat semicircular in cross-section towards pectoral-fin origin; length 2.63 (2.44–2.51) in pre-vent length; height 0.96 (0.76–0.78) times width. Snout elongate, narrowly triangular in lateral view, apex bluntly pointed; lateral prenarial margin angular; narrowly rounded in dorsal view; horizontal length 1.24 (1.04–1.23) times eye length, 0.93 (0.90–0.94) times interorbital space; horizontal prenarial length 2.50 (2.30–2.52) times in preoral length. Eye large, oval, length 4.13 (3.53–3.94) in head, 3.12 (3.10–4.48) times height; strongly notched posteriorly, notch not extending towards spiracle. Spiracle moderately-sized, somewhat triangular; no lobe-like fold on posterior margin; greatest diameter 3.30 (3.71–4.23) in eye length. Gill slits directed slightly anteroventrally from top to bottom; first four subequalin size,fifth longest, heightoffifth slit 2.5 (2.8–2.9)%TL.Mouthalmosttransverse,upperjawweaklyconcave,width 1.49 (1.43–1.63) in preoral length; upper labial

9Descriptions of new Australian chondrichthyans

Table 2. Proportional dimensions as percentages of total length for the holotype (CSIRO H 2625–06), 3 adult paratypes and2juvenileparatypesofCentrophorus westraliensis sp. nov.

Centrophorus westraliensis sp. nov. Holotype Adults Juveniles

Min. Max. Min. Max.Total length (mm) 909 774 866 371 414Precaudal length 81.8 78.8 81.1 77.9 78.3Pre-second dorsal length 64.9 63.8 64.9 60.9 62.0Pre-firstdorsallength 32.1 31.3 32.7 32.0 33.1Pre-vent length 61.1 58.1 61.7 56.5 58.2Prepelvic length 58.3 56.8 60.5 54.6 56.1Prepectoral length 22.5 22.9 23.7 25.6 26.4Head length 23.2 23.8 24.7 25.6 26.7Prebranchial length 19.3 20.2 21.2 22.5 22.8Prespiracular length 13.6 14.1 14.8 16.1 16.2Preorbital length 7.7 7.8 8.5 9.3 9.7Prenarial length 4.9 5.0 5.5 6.0 6.4Preoral length 11.3 10.9 12.4 13.7 14.2Inner nostril–labial furrow space 6.3 6.4 6.8 7.6 7.7Mouth width 7.6 7.6 7.8 7.6 10.4Upper labial furrow length 2.1 1.9 2.2 2.4 3.3Nostril width 1.6 1.8 1.9 2.1 2.2Internarial space 3.9 3.8 4.6 4.4 4.7Interorbital space 7.5 7.6 8.6 7.9 9.0Eye length 5.6 6.3 6.7 7.3 7.9Eye height 1.8 1.5 2.0 2.0 2.5Spiracle diameter - greatest 1.7 1.5 1.8 2.0 2.2First gill-slit height 2.2 1.8 1.8 2.2 2.3Fifth gill-slit height 2.5 2.8 2.9 2.3 2.9Interdorsal space 20.9 20.3 21.5 17.7 18.9Dorsal–caudal space 6.7 6.9 7.3 6.2 7.8Pectoral–pelvic space 32.7 30.9 31.8 25.4 27.8Pelvic–caudal space 16.1 13.7 14.6 13.6 14.6First dorsal length 18.4 17.1 18.2 16.1 18.5Firstdorsalsoftfinlength 12.6 11.3 11.9 10.1 11.7First dorsal anterior margin 11.2 11.4 12.3 11.5 13.2First dorsal base length 12.2 11.4 11.9 10.8 12.6First dorsal height 6.1 5.5 6.4 4.8 6.6First dorsal inner margin 6.1 5.8 6.5 5.5 6.0First dorsal posterior margin 10.2 8.8 9.7 8.1 9.8First dorsal spine length 2.7 2.4 2.5 1.4 2.1First dorsal spine base width 1.1 1.1 1.2 0.6 1.0Second dorsal length 14.5 13.5 13.6 14.1 14.2Seconddorsalsoftfinlength 8.5 8.4 8.9 8.0 8.5Second dorsal anterior margin 10.0 9.0 9.6 10.2 10.3Second dorsal base length 10.4 8.7 9.5 9.5 10.2Second dorsal height 4.6 4.6 5.0 4.5 4.6Second dorsal inner margin 4.2 4.6 4.9 4.8 4.8

10

Centrophorus westraliensis sp. nov. Holotype Adults Juveniles

Min. Max. Min. Max.Second dorsal posterior margin 7.4 7.1 7.6 6.3 7.6Second dorsal spine length 2.8 1.7 2.4 1.4 2.5Second dorsal spine base width 0.8 0.8 0.9 0.8 0.9Pectoral anterior margin 13.1 12.9 14.2 12.1 12.2Pectoral inner margin 13.4 12.5 14.9 11.3 11.6Pectoral base length 5.1 4.8 5.2 4.2 4.8Pelvic length 10.6 10.2 10.6 9.0 9.8Pelvic height 6.2 5.6 6.6 4.7 5.4Pelvic inner margin 5.5 5.4 6.1 4.8 5.5Dorsal caudal margin 17.4 18.5 20.2 21.5 22.2Preventral caudal margin 12.0 12.4 13.0 12.7 14.2Upper postventral caudal margin 6.9 6.1 6.6 7.9 8.6Lower postventral caudal margin 3.3 3.7 4.2 3.1 3.3Caudal fork width 6.5 6.5 6.9 6.9 7.1Caudal fork length 10.9 11.5 13.0 13.4 13.9Caudal terminal lobe 7.5 8.3 8.5 7.3 8.1Caudalsubterminalfinmargin 2.6 2.2 3.5 3.5 3.6Head width at anterior of nostrils 7.1 7.3 7.6 9.0 9.1Head width at mouth 8.7 8.9 9.7 10.7 11.3Head width 11.7 12.3 12.8 10.7 11.6Trunk width 9.2 8.9 10.2 7.2 8.5Abdomen width 10.0 8.7 11.0 6.9 9.0Tail width 5.4 5.2 5.3 4.3 4.9Caudal peduncle width 2.9 2.6 2.9 2.5 2.5Head height at mouth 5.6 5.5 5.9 6.0 6.3Head height 11.3 9.5 10.0 8.8 10.1Trunk height 13.1 9.6 11.4 9.2 11.7Abdomen height 12.9 11.6 12.7 9.6 12.8Tail height 7.0 6.4 6.8 6.1 7.3Caudal peduncle height 3.3 3.3 3.7 3.5 3.8Clasper outer length – – – 0.8 0.8Clasper inner length – – – 2.8 2.8Clasper base width – – – 0.5 0.5First dorsal midpoint–pectoral insertion 11.7 10.5 11.6 8.3 9.7First dorsal midpoint–pelvic origin 19.6 17.5 20.7 16.2 17.4Pelvicmidpoint–firstdorsalinsertion 16.0 15.3 17.2 12.9 13.8Pelvic midpoint–second dorsal origin 5.5 4.3 5.5 4.8 5.0

Table 2. cont’d.

furrows slightly longer than lower furrows; prominent postoral groove, more than twice length of upper labial furrows,extendingposterolaterally fromangleof jaws.Teeth strongly differentiated in upper and lower jaws;upper teeth small, cusps slightly oblique, relatively broad and triangular, bases sometimes overlapping; lower teeth much larger, cusps very strongly oblique, blade-like,

overlapping; sexual dimorphism was not established as there are no adult male specimens. Nostrils small, almost transverse; anterior nasal flap single lobed; internarialspace 2.88 (2.64–3.21) in preoral length, 2.51 (1.96–2.59) times nostril length. Dermal denticles (based on paratypeCSIROH2606–01)onflanksmall,flat,block-like, not overlapping; medial cusp blunt, no lateral cusps,

11Descriptions of new Australian chondrichthyans

several low blunt ridges.Denticles of juveniles (basedon CSIRO H 2358–01) are smaller, more upright, with acutely pointed crowns and more obvious ridges. First dorsal fin moderately small, strongly raked, broadlyrounded apically; anterior margin strongly convex; upper posterior margin straight, slanting well posteroventrally from top to bottom, moderately concave near free rear tip; free rear tip relatively thick basally, moderately long; inner margin of fin almost straight; insertion of baseextremelywellforwardofpelvic-finorigin,justposteriortofreereartipofpectoralfin;fin-spineoriginabovemidpectoral-fin inner margin; spine base broad, exposedanteriorly justbelow junctionof spineandsoftportionoffin;softportionoffinconnectedabovemid-pointoftotal spine length; spine rapidly tapering distally (tip often damaged), anterior margin almost straight; exposed portion of spine sloping strongly posterodorsally from base to apex, subequal in length to exposed portion of second dorsal-fin spine; pre-first dorsal-fin length 3.11(3.06–3.19) times in TL; first dorsal-fin length 3.04(2.83–3.11) times its height, 1.27 (1.26–1.34) times second dorsal-fin length; first dorsal-fin height 1.33(1.20–1.30) times second dorsal-fin height; exposedfirstdorsalspinelength0.45(0.39–0.45)timesheightoffin. Second dorsal fin ofmoderate size, slightly raked;anterior margin very slightly convex, apex moderately rounded; posterior margin very weakly concave, sloping strongly posteroventrally from apex; free rear tip greatly elongated, inner margin length 0.92 (0.95–1.00) times fin height; second dorsal-fin length 3.19 (2.69–2.98)times its height; spine length 0.61 (0.38–0.49) in height of fin; fin-spine origin just posterior to free rear tip ofpelvicfin,exposedjustbelowlevelofjunctionwithspineandsoftportionoffin; seconddorsal spinemoderatelybroad based, tapering rapidly distally, sharply pointed; interdorsal space 1.08 (1.07–1.17) in prepectoral length, 1.54 (1.50–1.60) in pre-first dorsal length; interdorsalgrooveweak.Pectoralfinlarge,anteriormarginweaklyconvex; inner margin weakly convex anteriorly, almost

Figure 6. Ventral view of the head of Centrophorus westraliensis sp. nov., preserved holotype (CSIRO H 2625–06, female 909 mm TL).

straight posteriorly, length 13.4 (12.5–14.9)% TL; apex moderately rounded, lobe-like but not falcate; posterior margin almost straight from apex to free rear tip; free rear tip greatly elongated, extending to just posteriorto midpoint of first dorsal-fin base; base very short,2.58 (2.59–2.78) in anterior margin length. Pelvic fins moderately large, anterior margin almost straight,posterior margin very weakly concave, apex moderately rounded, free rear tip acute. Caudal peduncle moderately long, tapering slightly towards caudal fin; subcircularin cross-section anteriorly, becoming more compressed posteriorly; ventral groove weak (better developed in some paratypes); no lateral keels; pelvic–caudal space 2.04 (2.23–2.32) in pectoral–pelvic space, 1.40 (1.62–1.71) in prepectoral length; dorsal–caudal space 3.10 (2.83–3.04) in interdorsal length; precaudal pits absent. Caudalfinrelativelylong,postventralmarginmoderatelyconcave, terminal lobe moderately large; apex of lower lobe moderately rounded; dorsal caudal margin 1.34 (1.18–1.34) in head length; length of lower caudal lobe 1.45 (1.44–1.56) in upper lobe length. Total vertebral centra 117 (112–117), monospondylous precaudal centra 57 (55 or 56), diplospondylous precaudal centra 30 (29–33), precaudal centra 87 (85–88) and diplospondylous caudal centra 30 (27–29). Teeth count: 38/29 (holotype).

COLOUR.— Fresh specimens: (based on paratype CSIRO H 2606–01): dorsal surfaces uniformly light greyish to brownish; ventral surfaces much paler, with ventral surface of head somewhat darker than belly area. Dorsalfinswithafaintdarkblotchextendingfrommidanteriormargin tonearapex tomidportionofsoftfin;distinctwhitishposteriormargin;finspinespale,greyishbrown.Caudalfinmostlygreyish,withadiffuse,narrowwhitish posterior margin. Injuveniles (based on CSIRO H 2358–01): similar coloration to adult, but ventral surfaces less demarcated from dorsal surfaces.Dorsal finswithvery distinct, blackish oblique blotch from lower anterior margintofininsertion;distinctwhiteblotchonposterior

Figure 7.CuspsoftheflankdenticlesofCentrophorus westraliensis sp. nov. (holotype CSIRO H 2625–06, female 909 mm TL).

12

marginfromfinapextomidposteriormargin.Pectoralfinwith dark terminal blotch and a narrow, whitish posterior margin.Caudalfinwithverydistinctivewhitishmargin,quite broad on terminal lobe; distinct blackish fringe along mostofdorsalcaudalmargintonearfinapex,becomingablackish blotch over most of the terminal lobe; lower soft fin areamostly dark.Preserved specimens: (based on holotype): similar in coloration; deciduous denticles over bodyandheadgivingappearanceofsmallwhiteflecks(alsoevidentinotheradulttypespecimens).Dorsalfinsmostly uniform in coloration, slightly darker near apex, whiteposteriormarginbarelyevident.Caudalfinmostlyuniformincoloration.Pectoralandpelvicfinswithpaleposterior margins.

SIZE.— Type series consists of five females between414 and 909 mm TL and a single immature male of 371 mm TL.

DISTRIBUTION.— Type specimens collected from the mid-continental slope of Western Australia from south ofCapeLeeuwin(35°02′S)toSharkBay(27°05′S)indepths of 616–750 m (Fig. 11).

ETYMOLOGY.— Named in allusion to the known geographic range of this species (Western Australia).

VERNACULAR.— We propose the official Englishcommon name of “Western Gulper Shark” in allusion to its geographic distribution in Australia.

REMARKS.— Centrophorus westraliensis was previously considered to be conspecific with C. harrissoni, but Last & Stevens (1994) noted that eastern and western populations need to be critically compared. This new species is similar to other members of the longnose Centrophorus subgroup, in particular C. harrissoni. Adults of C. westraliensis differ from those of C. harrissoni inhavingdorsalfinsslightlyfurtherapart(interdorsal space 20.3–21.5 vs. 18.2–20.6% TL, 2.8–3.1 vs. 2.2–2.6 times dorsal–caudal space); a longer pectoral anterior margin (12.9–14.2 vs. 11.9–12.6% TL); smaller firstdorsalfin(baselength11.4–12.2vs.12.7–14.1%TL,height 5.5–6.4 vs. 6.2–7.0% TL); a smaller mouth (width 7.6–7.8 vs. 7.8–8.5% TL, 3.1–3.2 vs. 2.7–3.0 in head length);alargermid-firstdorsalbasetopectoralinsertion(10.5–11.7 vs. 8.4–9.7% TL) and a narrower head (head width 11.7–12.8 vs. 13.2–14.0% TL, head width at anterior of mouth 8.7–9.7 vs. 10.2–10.7% TL, 3.2–3.7 vs. 2.8–3.0 times pre-first dorsal length). Juveniles of C. westraliensis differ from those of C. harrissoni in the following measurements and ratios: pre-pelvic length 54.6–56.1 vs. 52.6–53.8% TL; pre-vent length 56.5–58.2 vs. 55.4–55.6% TL; upper labial furrow length 2.4–3.3 vs.2.0–2.5%TL;midfirstdorsal-finbase topelvic-finorigin space 16.2–17.4 vs. 13.4–15.4% TL.

Adults of Centrophorus westraliensis differ from those of C. isodon in having a longer, slightly broader head (head

length 23.2–24.7 vs. 22.0–23.3% TL; head width 11.7–12.8 vs. 9.7–11.9% TL, 5.1–5.5 vs. 5.7–6.9 in pre-second dorsal length); shortermiddorsal-finbase topelvic-finorigin(17.5–20.7vs.21.9–23.3%TL);dorsalfinsclosertogether (interdorsal space 20.3–21.5 vs. 22.4–25.1% TL, 2.8–3.1 vs. 3.4–4.3 times dorsal–caudal space) and upper teeth with much more oblique cusps. Juveniles of C. westraliensis differ from those of C. isodon in the following measurements and ratios: pre-second dorsal length 60.9–62.0 vs. 63.0–64.3% TL; pectoral–pelvic space 25.4–27.8 vs. 29.1–29.5% TL; preoral length 13.7–14.2 vs. 11.0–11.9% TL; inner nostril to upper labial furrow 7.6–7.7 vs. 6.0–6.5% TL; head width at anterior of nostrils 9.0–9.1 vs. 8.2–8.3% TL; preorbital length 9.3–9.7 vs. 8.1–8.3% TL, 6.6–7.2 vs. 8.1–8.5 in pre-second dorsal length; first dorsal-fin soft length10.1–11.7vs.12.2–13.0%TL;midfirstdorsal-finbasetopelvic-finoriginspace16.2–17.4vs.18.8–19.5%TL.

Adults of this species differ from C. tessellatus in having a longer horizontal prenarial length (4.5–5.9 vs. 3.7% TL); a longer pectoral–pelvic space (30.9–32.7 vs. 27.9% TL); a shorter pelvic–caudal space (13.7–16.1 vs. 17.9% TL);ashorterfirstdorsalfin(length17.1–18.4vs.20.1%TL) and no white borders on the gill slits.

Females of C. westraliensis have similar upper teeth to the holotype of C. seychellorum, but these two species differ in horizontal prenarial length (4.5–4.9 vs. 2.2% TL);seconddorsal-finheight(4.6–5.0vs.3.7%TL)andpre-pectoral length (22.5–23.7 vs. 20.9% TL).

Centrophorus zeehaani sp. nov.

Figs 8–10, Table 3

Centrophorus uyato (in part): Last & Stevens, 1994: pp 52, 60, 61,keyfig.57,fig.8.5,pl.4;Daleyet al., 2002: pp 53; Gomon et al.1994:pp92,94,figs30,31;Yearsleyet al., 2001: pp 35, 360.

Holotype. CSIRO H 6628–05, adult male 893 mm TL, south-west of Coffin Bay, South Australia, 35°14′ S,134°29′E,360–600m,28July2005.Paratypes. 8 whole specimens: AMS I 44310–001, adult male 826 mm TL, CSIRO H 6628–01, immature male 506 mm TL, CSIRO H 6628–02, immature male 645 mm TL, CSIRO H 6628–03, adult male 875 mm TL, CSIRO H 6628–04, adult male 910 mm TL, CSIRO H 6628–06, adult male 852 mm TL, CSIRO H 6628–07, adult male 906 mm TL, NMV A 29736–001, adult male 820 mm TL, collected with holotype.Other material. 5 whole specimens: CSIRO CA 4104, adult male 843 mm TL, east of Gabo Island, Victoria, 37°40′S,150°15′E,504–508m,04May1984;CSIROH 866–02, immature male 456 mm TL, CSIRO H 867–01, female 439 mm TL, east of Jervis Bay, New South Wales,34°58′S,151°09′E,490–576m,10Sep1986;

13Descriptions of new Australian chondrichthyans

CSIRO H 2268–02, adult male 800 mm TL, west of Bunbury,WesternAustralia,33°03′S,114°25′E,701m,10 Feb 1989; CSIRO H 6504–05, adult male 861 mm TL, eastofJervisBay,NewSouthWales,35°12′S,150°58′E, 320–500 m, July to Aug 2003. 8 skeletal specimens: CSIRO H 6307–01, female 1027 mm TL, east of Flinders Island, Tasmania, ca. 40° S, 149° E, 350–430 m, 12 July 2004; CSIRO H 6309–01, adult male 865 mm TL, east of Flinders Island, Tasmania, ca. 40° S, 149° E, 400– 450 m, 01 Aug 2004; CSIRO H 6503–01, adult male 872 mm TL, CSIRO H 6503–02, female 991 mm TL, CSIRO H 6503–03, female 1023 mm TL, CSIRO H 6503–04, female 987 mm TL, CSIRO H 6503–05, female 957 mm TL, CSIRO H 6503–06, adult male 867 mm TL, north-east of Flinders Island, Tasmania, 39°20′S,148°45′E,370–420m,07Apr2003.

DIAGNOSIS.— A moderate-sized species of Centrophorus with the following combination of adult characters: pre-second dorsal length 62.8–64.8% TL, 8.3–9.8timesdorsal–caudalspace;pre-firstdorsallength28.3–30.7% TL; interdorsal space 20.0–23.7% TL, 3.0–3.5 times dorsal–caudal space; dorsal–caudal space 6.4–7.9% TL, 4.2–4.7 in pectoral–pelvic space; head long and moderately robust (length 23.3–24.3% TL, 2.5–2.9 times mouth width; width 11.8–13.1% TL, 4.9–5.4 in pre-second dorsal length; width at anterior of nostrils 5.5–6.3% TL); snout moderately long (preoral length 9.4–10.1% TL, 1.6–1.9 times head height at anterior of mouth, 1.0–1.2 times mouth width; horizontal preorbital length 5.6–6.3% TL; horizontal prenarial length 3.4–4.2% TL); mouth large (width 8.3–9.6% TL); pectoral fin moderately sized (anterior margin 11.4–12.6% TL,2.3–2.8timesbaselength);caudalfinlarge(dorsalcaudal

margin 17.9–20.8% TL; 2.6–2.9 times dorsal–caudal space);firstdorsalfinmoderately-sized(height6.0–7.0%TL), spine relatively robust (base width 0.9–1.2% TL). Dorsalfinsof juvenileswithadistinctblackishmarginextending from mid-anterior margin to near free rear tip; adults with a less distinct, but obvious, dark margin. Upper teeth of adults of both sexes upright, only slightly oblique laterally; different shape and much smaller than lower teeth;thoseofjuvenileswithmoreobliquecusps,butnotstrongly oblique. Tooth row count (based on non-type skeletal specimens, n=8): 37–45/30–33. Flank denticles flat, block-like,notoverlapping, scallopededges.Totalvertebral centra 114–117 (mean 115.4), monospondylous precaudal centra 54–56 (55.2), diplospondylous precaudal centra 28–31 (29.4), precaudal centra 84–86 (84.7) and diplospondylous caudal centra 31 (30–32).

DESCRIPTION.— Body fusiform, moderately elongate, nape moderately humped; deepest near firstdorsal-finspine,trunkheight1.08(0.97–1.36in6adultparatypes) times width, 1.01 (0.92–0.19) times abdomen height; head moderately elongate, length 23.3 (23.3–24.3)% TL; caudal peduncle moderately slender, 13.7 (12.0–14.8)% TL. Head moderately robust, broad, width 1.10 (1.04–1.19) times trunk width, 1.07 (1.03–1.45) times abdomen width; depressed forward of spiracles, becoming somewhat semicircular in cross-section towards pectoral-fin origin; length 2.59 (2.26–2.58) inpre-vent length; height 0.88 (0.83–0.96) times width. Snout relatively short, narrowly rounded in lateral view, apex bluntly pointed; lateral prenarial margin angular; narrowly rounded in dorsal view; horizontal length 0.97 (0.88–1.12) times eye length, 0.73 (0.69–0.82) times interorbital space; horizontal prenarial length

Figure 8. Lateral view of Centrophorus zeehaani sp. nov.: A. preserved holotype (CSIRO H 6628–05, adult male 893 mm TL); B. preserved paratype (CSIRO H 6628–01, immature male 506 mm TL).

A

B

14

Table 3. Proportional dimensions as percentages of total length for the holotype (CSIRO H 6628–05), 6 adult paratypes and2juvenileparatypesofCentrophorus zeehaani sp. nov.

Centrophorus zeehaani sp. nov. Holotype Adults Juveniles

Min. Max. Min. Max.Total length (mm) 893 820 910 506 645Precaudal length 79.7 78.7 81.9 78.1 79.6Pre-second dorsal length 63.6 62.8 64.8 62.1 63.1Pre-firstdorsallength 30.7 28.3 30.3 30.1 30.3Pre-vent length 60.4 57.8 61.0 57.5 58.6Prepelvic length 58.2 55.8 58.3 55.9 56.6Prepectoral length 22.2 21.8 22.8 24.5 24.9Head length 23.3 23.3 24.3 25.4 25.5Prebranchial length 19.3 19.4 20.1 21.8 21.9Prespiracular length 12.1 12.2 13.5 13.9 14.6Preorbital length 6.1 6.3 6.8 7.2 7.4Prenarial length 4.2 4.1 4.5 5.0 5.0Preoral length 9.4 9.5 10.1 10.8 11.3Inner nostril–labial furrow space 5.9 6.1 6.6 6.8 6.9Mouth width 8.9 8.3 9.6 9.3 9.3Upper labial furrow length 2.0 1.5 2.2 2.0 2.3Nostril width 1.6 1.6 1.9 1.9 2.1Internarial space 3.6 3.3 3.5 3.7 3.9Interorbital space 7.6 7.7 8.6 7.9 8.2Eye length 5.8 5.4 6.5 6.5 7.3Eye height 2.2 1.7 2.2 2.1 2.4Spiracle diameter - greatest 2.1 1.6 1.9 1.6 2.1First gill-slit height 2.8 2.2 2.6 2.7 2.8Fifth gill-slit height 3.6 3.0 3.8 3.5 3.6Interdorsal space 21.9 20.0 23.7 21.5 22.4Dorsal–caudal space 6.9 6.4 7.9 7.7 7.7Pectoral–pelvic space 32.4 30.0 34.1 29.4 29.6Pelvic–caudal space 13.7 12.0 14.8 12.4 14.2First dorsal length 17.8 18.5 19.9 17.8 18.3Firstdorsalsoftfinlength 11.8 11.1 12.7 11.5 12.0First dorsal anterior margin 10.9 12.0 13.7 13.0 13.2First dorsal base length 11.8 12.2 13.8 11.0 12.7First dorsal height 6.0 6.1 7.0 6.8 7.1First dorsal inner margin 6.2 6.2 7.0 5.8 6.7First dorsal posterior margin 9.5 9.0 11.0 9.4 10.6First dorsal spine length 2.6 2.1 3.1 3.0 3.2First dorsal spine base width 0.9 0.9 1.2 1.1 1.2Second dorsal length 14.2 12.8 14.5 13.4 13.4Seconddorsalsoftfinlength 8.3 7.4 8.8 8.2 8.5Second dorsal anterior margin 9.4 9.5 10.5 10.0 10.0Second dorsal base length 9.6 9.2 10.0 8.6 8.8Second dorsal height 4.7 4.7 5.3 5.2 5.2Second dorsal inner margin 4.5 3.6 4.9 4.5 5.0

15Descriptions of new Australian chondrichthyans

Centrophorus zeehaani sp. nov.

Holotype Adults Juveniles Min. Max. Min. Max.

Second dorsal posterior margin 7.2 6.0 7.5 6.6 6.9Second dorsal spine length 2.2 2.4 3.3 3.4 3.5Second dorsal spine base width 0.9 0.7 1.0 1.0 1.0Pectoral anterior margin 11.4 11.9 12.6 12.6 13.0Pectoral inner margin 13.6 12.3 14.6 13.0 13.0Pectoral base length 4.8 4.5 5.2 4.5 5.2Pelvic length 11.2 11.1 12.2 10.7 11.0Pelvic height 6.5 6.0 6.7 5.0 5.4Pelvic inner margin 6.7 6.0 7.0 5.1 6.1Dorsal caudal margin 19.6 17.9 20.8 19.8 21.7Preventral caudal margin 12.5 12.2 13.8 13.0 14.4Upper postventral caudal margin 7.7 7.2 8.5 8.0 8.8Lower postventral caudal margin 4.1 4.6 5.6 4.1 4.5Caudal fork width 7.1 7.2 7.8 7.6 7.8Caudal fork length 11.8 11.8 12.7 11.9 14.1Caudal terminal lobe 8.8 8.3 9.7 9.1 9.9Caudalsubterminalfinmargin 3.4 2.7 3.1 1.7 1.7Head width at anterior of nostrils 5.5 5.7 6.3 6.1 6.8Head width at mouth 9.9 9.3 10.4 10.9 11.2Head width 12.6 11.8 13.1 12.2 12.7Trunk width 11.4 10.0 12.1 10.5 11.2Abdomen width 11.7 10.4 12.7 8.4 9.3Tail width 5.8 5.1 5.8 4.8 4.9Caudal peduncle width 3.0 2.8 3.4 2.8 2.8Head height at mouth 5.8 5.4 6.5 5.3 6.3Head height 11.0 10.9 12.1 10.5 10.7Trunk height 12.2 12.3 14.7 10.9 11.3Abdomen height 12.1 10.8 15.2 10.3 11.0Tail height 6.8 6.4 7.2 6.4 7.3Caudal peduncle height 3.8 3.7 4.2 3.8 3.9Clasper outer length 3.7 3.4 4.0 – –Clasper inner length 8.3 7.1 8.5 – –Clasper base width 1.1 1.0 1.2 – –First dorsal midpoint–pectoral insertion 8.2 7.4 9.6 7.1 7.5First dorsal midpoint–pelvic origin 22.0 19.3 22.7 20.1 21.1Pelvicmidpoint–firstdorsalinsertion 17.9 14.5 18.7 16.7 17.5Pelvic midpoint–second dorsal origin 3.8 3.8 5.1 4.4 4.5

Table 3. cont’d.

2.49 (2.30–2.79) times in preoral length. Eye large, oval, length 4.05 (3.49–4.37) in head, 2.68 (2.62–3.58) times height; strongly notched posteriorly, notch not extending towards spiracle. Spiracle moderately-sized, semicircular; no lobe-like fold on posterior margin; greatest diameter 2.79 (2.99–4.49) in eye length. Gill slits directed slightly anteroventrally from top to bottom;

first four subequal in size, fifth longest, height of fifthslit 3.6 (3.0–3.8)% TL. Mouth almost transverse, upper jawslightlyconcave,width1.05(1.02–1.22) inpreorallength; upper labial furrows subequal to or slightly longer than lower furrows; prominent postoral groove, more than twice length of upper labial furrows, extending posterolaterallyfromangleofjaws.Teethofadultmales

16

strongly differentiated in upper and lower jaws; upperteeth small, cusps upright, narrow and triangular, bases not usually overlapping; lower teeth much larger, cusps very strongly oblique, blade-like, overlapping, apices often somewhat upright, recurved; sexual dimorphism was not established as there are no adult female type specimens. Nostrils small, almost transverse; anterior nasal flapformed as a large subtriangular lobe with a somewhat rudimentary secondary lobe mesially; internarial space 2.62 (2.77–3.10) in preoral length, 2.19 (1.74–2.21) times nostril length. Dermal denticles (based on holotype CSIROH6628–05)onflanksmall,flat,block-like,notoverlapping; medial cusp blunt, no lateral cusps, ridges indistinct. Denticles of juveniles (based on CSIRO H6628–01) smaller, more upright, with more pointed crowns and ridges more obvious. First dorsal fin moderatelysmall, raked, broadly rounded apically (somewhat broadly angular in CSIRO H 6628–01); anterior margin moderately convex; upper posterior margin straight to slightly convex, slanting well posteroventrally from top to bottom, moderately concave near free rear tip; free rear tip relatively thick basally, moderately long; inner marginoffinalmoststraight(weaklyconcaveinCSIROH 6628–01); insertion of base extremely well forward of pelvic-finorigin,posteriortofreereartipofpectoralfin(overfreerear tip inCSIROH6628–01,02);fin-spineorigin abovemid pectoral-fin innermargin; spine basebroad, exposed anteriorly just below junction of spineandsoftportionoffin;softportionoffinconnectedaboutlevel of two thirds of total spine length; spine rapidly tapering distally when not damaged, anterior margin almost straight to weakly convex; exposed portion of spine sloping strongly posterodorsally from base to apex, usually shorter than exposed portion of second dorsal-fin spine; pre-first dorsal-fin length 3.26 (3.30–3.54)times in TL; first dorsal-fin length 2.98 (2.58–3.14)times its height, 1.25 (1.33–1.51) times second dorsal-finlength;firstdorsal-finheight1.26(1.19–1.39)timesseconddorsal-finheight;exposedfirstdorsalspinelength0.44 (0.31–0.47) timesheight offin.Seconddorsalfinmoderately small, slightly raked; anterior margin slightly convex, apex moderately rounded (narrowly rounded in CSIRO H 6628–01, 02); posterior margin very weakly concave (moderately concave in CSIRO H 6628–01, 02), sloping strongly posteroventrally from apex; free rear tip greatly elongated, inner margin length 0.96 (0.73–0.96) times fin height; second dorsal-fin length 3.01 (2.56–2.89) times its height; spine length 0.47 (0.47–0.67) in heightoffin;fin-spineoriginanteriortofreereartipofpelvicfin,exposedjustbelowlevelofjunctionwithspineandsoftportionoffin; seconddorsal spinemoderatelybroad based, tapering distally, sharply pointed when undamaged; interdorsal space 1.01 (0.94–1.14) in prepectoral length, 1.40 (1.22–1.49) in pre-first dorsallength;interdorsalgrooveweak.Pectoralfinmoderatelylarge, anterior margin weakly convex; inner margin weakly convex anteriorly, almost straight posteriorly, length 13.6 (12.3–14.96)% TL; apex moderately rounded to somewhat angular, lobe-like but not falcate; posterior

margin almost straight from apex to free rear tip; free reartipgreatlyelongated,extendingtojustposteriortomidpointoffirstdorsal-finbase(extendingtofirstdorsal-fininsertioninCSIROH6628–01,02);baseveryshort,2.40 (2.33–2.77) in anteriormargin length. Pelvic finsmoderately large, anterior margin almost straight (weakly convex in CSIRO H 6628–02), posterior margin weakly concave to nearly straight, apex moderately rounded, free rear tip acute to narrowly rounded. Caudal peduncle moderately long, compressed, tapering slightly towards caudalfin;ventralgrooveweak;nolateralkeels;pelvic–caudal space 2.36 (2.09–2.63) in pectoral–pelvic space, 1.61 (1.50–1.98) in prepectoral length; dorsal–caudal space 3.17 (2.79–3.46) in interdorsal length; precaudal pitsabsent.Caudalfinrelativelylong,postventralmarginmoderately concave (nearly straight in CSIRO H 6628–01), terminal lobe moderately large, deep (relatively shallow in CSIRO H 6628–01, 02); apex of lower lobe narrowly to moderately rounded; dorsal caudal margin 1.19 (1.12–1.32) in head length; length of lower caudal lobe 1.57 (1.47–1.58) in upper lobe length. Total vertebral centra 115 (114–117), monospondylous precaudal centra 56 (54–56), diplospondylous precaudal centra 28 (28–31), precaudal centra 84 (84–86) and diplospondylous caudal centra 31 (30–32). Tooth row count (based on non-type skeletal specimens): 37–45 (mean = 40; n=8)/30–33 (mean = 31.7, n=7).

COLOUR.— Preserved specimens: (based on adult male paratypes) Upper surface of body uniformly light brownish with a distinct pale reddish tinge, becoming paler ventrally; ventral surfaces much paler; light and dark tonal areas not well demarcated on body with light and dark areas almost blending together; light and dark tonal areas well demarcated on head, extending justbelowlowermarginofeyeandjustbelowuppermarginofgill slits,pectoral-finoriginpale;eye–spiraclespacewith a distinct pale whitish blotch anteriorly, less distinct near spiracle (less obvious in some paratypes). Dorsal fins pale with a distinct, broad, diffuse-edged, duskymarginextendingfromaboutleveloffin-spineapicestojustposteriorofthemaximumconcavityoftheposteriormargin;finspinesdarkbrownish.Pectoralandpelvicfinupper surfaces similar in colouration to dorsal surface but darker distally, with narrow whitish posterior margins. Caudalfinmostlygreyish;postventralmarginwithaverynarrow darkish postventral border (indistinct in some paratypes), a narrow whitish submarginal bar, demarcated by a broad dusky, diffuse-edged marking for its entire length;terminallobedarkerthanrestoffinwithanarrowwhitish terminal margin. Juveniles: (based on CSIRO H 6628–01, 02) upper surface of body distinctly medium greyish, lacking a reddish tinge (CSIRO H 6628–02 with a number of deciduous denticles resembling small white flecks);adistinctdarkgreyishblotchoverupperportionof gill slits, extending dorsally to level of upper margin of spiracle (less distinct in CSIRO H 6628–02). Dorsal finsmuchmoredistinct,broad,blackishmargins,insamelocationasonadults;freereartipswhitish;finspinebases

17Descriptions of new Australian chondrichthyans

whitish.Pectoralfinuppersurfacewithamoredistinctdark greyish distal marking and white posterior margin. Pelvicfinpalewithwhitishposteriormargin.Caudalfinmostly similar to that of adults but with a much more distinct darker terminal lobe and apical half of preventral margin with a very narrow, blackish border.

SIZE.— Type series consists of 7 adult males between 820 and 910 mm TL and two immature males of 506 and 645 mm TL. Largest female examined was 1027 mm TL (CSIRO H 6307–01) and smallest free-swimming individual was 439 mm TL (CSIRO H 867–01).

DISTRIBUTION.— Known from the continental slope of southern Australia from off Forster in New South Wales (ca. 32° S; K. Graham, pers. comm.) to offBunbury inWesternAustralia (33°03′S;CSIROH2268–02), including Tasmania, in depths of 208–701 m, but usually found in depths >400 m (Fig. 11).

ETYMOLOGY.— Named after the commercial vessel Zeehaan fromwhichthefirstspecimensof thisspeciesfrom Tasmanian waters were collected in 1979 during the firstsurveytomapandexploreunchartedgrounds(seeLast & Harris, 1981).

VERNACULAR.—TheofficialEnglishcommonnameof“SouthernDogfish”usedbyLast&Stevens(1994)forC. uyato in allusion to the southern Australian distribution of this species is adopted for this species.

REMARKS.— Centrophorus zeehaani was previously considered to be C. uyato (Last & Stevens, 1994; Gomon et al., 1994), but Garman’s (1906) placement of Rafinesque’s (1810: 3–14, pl. 14, fig. 2) Squalus uyatus into the genus Centrophorus was incorrect. The

Figure 9. Ventral view of the head of Centrophorus zeehaani sp. nov. holotype (CSIRO H 6628–05, adult male 893 mm TL).

illustration of S. uyatus clearly depicts a Squalus species and not a Centrophorus species (as also highlighted by Muñoz-Chápuli & Ramos, 1989), which is particularly evidentfromthelackofaterminallobeonthecaudalfintypical of members of the latter genus. This new species issuperficiallysimilartoothermembersofthelongnoseCentrophorussubgroup,andiscommonlymis-identifiedas C. harrissoni in areas where these two species co-occur. Adults of C. zeehaani differ from C. tessellatus in having a shorter horizontal head length (22.0–23.3 vs. 24.8% TL); a shallower head (10.9–12.1 vs. 9.9% TL); a narrower mouth (8.3–9.6 vs. 7.4% TL); pectoral and pelvic fins closer together (pectoral–pelvic space30.0–34.1 vs. 27.9% TL); a shorter caudal peduncle (pelvic–caudal space 12.0–14.8 vs. 17.9% TL) and gill slits mostly pale but with no white borders vs. darker with white borders.

Adults of C. zeehaani differ from those of C. isodon in having a slightly longer, broader head (head length 23.3–24.3 vs. 22.0–23.3% TL; head width 11.8–13.1 vs. 9.7–11.9% TL, 4.9–5.4 vs. 5.7–6.9 in pre-second dorsal length); a longer pre-second dorsal length (62.8–64.8 vs.66.2–68.0%TL);largergillslits(firstgillslitheight2.2–2.8 vs. 1.7–1.9% TL, 2.0–2.9 vs. 3.5–4.1 in head width at anterior of nostrils); more robust dorsal spines (exposedfirstdorsal-spinebasewidth0.9–1.2vs.0.7%TL);alongerpelvicfin(pelvic-finlength11.1–12.2vs.7.6–10.4% TL); a deeper head (head height 10.9–12.1 vs. 8.1–9.5% TL); a wider mouth (mouth width 8.3–9.6 vs. 7.4–7.9% TL, 1.0–1.2 vs. 1.3–1.5 in preoral length) and dorsal fins blackish apically without a white posteriormargin vs. uniformly darkish with a sometimes indistinct white posterior margin. Juveniles of C. zeehaani differ from those of C. isodon in the following measurements and ratios: a longer, broader head (head length 25.4–25.5

Figure 10.CuspsoftheflankdenticlesofCentrophorus zeehaani sp. nov. (holotype CSIRO H 6628–05, adult male 893 mm TL).

18

vs. 24.2–24.8% TL; head width 12.2–12.7 vs. 9.2–11.2% TL); a shorter, narrower snout (preorbital length 7.2–7.4 vs. 8.1–8.3% TL, width at anterior of nostrils 6.1–6.8 vs. 8.2–8.3% TL); more robust dorsal spines; larger gill slits (firstgillslitheight2.7–2.8vs.2.1%TL);adeeperhead(head height 10.5–10.7 vs. 7.1–8.7% TL); a wider mouth (mouthwidth9.3vs.8.1–8.5%TL)anddorsalfinswithdistinct, broad blackish apices and posterior margins vs. dorsalfinswithablackishanteriorblotchextendingoverapex and a white posterior blotch.

Adult males of C. zeehaani have similar, upright upper teeth to the adult male holotype of C. seychellorum (Baranes, 2003), but these two species differ in pelvic–caudal space (12.0–14.8 vs. 19.1% TL); pectoral anterior margin(11.4–12.6vs.13.4%TL);seconddorsal-finheight(4.7–5.3 vs. 3.7% TL); mouth width (8.3–8.6 vs. 7.1% TL) and pre-pectoral length (21.8–22.8 vs. 20.9% TL).

Adults of C. zeehaani differ from those of C. harrissoni and C. westraliensis in having a shorter, narrower snout (preorbital length 6.1–6.8 vs. 7.5–8.6% TL, 10.0–11.6 vs. 7.7–9.3 times pre-second dorsal length, prenarial length 4.1–4.5 vs. 4.9–5.8% TL, width at anterior of nostrils 5.5–6.3 vs. 7.1–7.9% TL); a shorter preoral length (9.4–10.1 vs. 10.9–12.4% TL); a slightly longer pelvic fin(length 11.1–12.2 vs. 9.9–11.2% TL); a short internarial space (3.3–3.6 vs. 3.8–4.6% TL); height of head at anterior of mouth 1.6–1.9 vs. 2.0–2.2 in preoral length and dorsal fins with a broad, dusky to blackish apical

marking(usuallyobviouswhenfresh)vs.dorsalfinswitha narrow white posterior margin (sometimes indistinct in larger specimens). Juveniles of C. zeehaani differ from those of C. harrissoni and C. westraliensis in having dorsalfinswithadistinct,broadblackishapicalmarkingsand pale free rear tips vs. a blackish anterior blotch and white posterior blotch. They also differ in the following measurements and ratios: prespiracular length 13.9–14.6 vs. 15.9–16.2% TL; preorbital length 7.2–7.4 vs. 9.3–9.7% TL; interdorsal space 21.5–22.4 vs. 16.1–19.7% TL; pectoral–pelvic space 29.4–29.6 vs. 25.0–27.8% TL; preorallength10.8–11.3vs.13.0–14.2%TL;pectoral-fininnermarginlength13.0vs.10.6–11.6%TL;firstdorsalspine-length 3.0–3.2 vs. 1.4–2.1% TL; second dorsal spine-length 3.4–3.5 vs. 1.4–2.5% TL; width at anterior of nostrils6.1–6.8vs.8.6–9.2%TL;caudal-finsubterminalmargin1.7vs.2.9–3.7%TL;midpelvic-finbasetofirstdorsal-fininsertion16.7–17.5vs.11.0–13.8%TL.

The upper teeth of adults of Centrophorus zeehaani are not strongly sexually dimorphic (based on skeletal specimens) as they are in C. harrissoni and C. seychellorum, in which the upper teeth of females have strongly oblique cusps and those of males have upright, pointed cusps. Those of C. isodon also show sexual dimorphism but with the cusps of female upper teeth less strongly oblique than in the above two species. The larger female types of C. westraliensis have similar-shaped upper teeth to C. isodon, and not as strongly oblique as C. harrissoni or C. seychellorum. The upper teeth of the adult

Figure 11. Map showing the geographic distribution (not indicative of depth ranges) of the three longnose Centrophorus species, C. harrissoni (dark grey), C. westraliensis (pale grey) and C. zeehaani (black), which occur in Australian waters. Collection localities of the holotype of each of the species are denoted by stars.

19Descriptions of new Australian chondrichthyans

male paratypes and of female material examined (CSIRO H 6307–01, CSIRO H 6503–02, 03, 04, 05) all have upright cusps becoming only very slightly oblique laterally.

The biological data which has been collected for this species so far has revealed that females give birth to a single pup (K. Graham, NSW Department of Primary Industries and R. Daley, CSIRO Marine & Atmospheric Research, unpubl. data). In contrast, all pregnant females of C. isodon examined in Indonesia contained 2 embryos (unpubl. data) and most pregnant females of C. harrissoni examined from southeastern Australia contained two pups (K. Graham and R. Daley, unpubl. data). This particularly low fecundity highlights the vulnerability of members of thisgenustofishingpressure.

Comparative material. Centrophorus isodon: CSIRO H 5857–01, immature male 540mmTL,Kedongananfishmarket,Bali, Indonesia,9 Apr 2001; CSIRO H 5889–15, female 530 mm TL, Kedonganan fish market, Bali, Indonesia, July 2002;CSIROH5875–04,female989mmTL,TanjungLuarfishlanding site, Lombok, Indonesia, 26 Mar 2002; CSIRO H 6125–02,female965mmTL,Kedongananfishmarket,Bali, Indonesia, 27 Aug 2002; CSIRO H 6138–01, female 952mmTL, Tanjung Luar fish landing site, Lombok,Indonesia, 25 Mar 2002; CSIRO H 6233–02, immature male 306 mm TL, CSIRO H 6233–03, immature male 304 mm TL, CSIRO H 6233–04, immature male 305 mm TL, CSIRO H 6233–05, female 241 mm TL, CSIRO H 6233–06,immaturemale256mmTL,Kedongananfishmarket, Bali, Indonesia, 16 Mar 2005; SAM unregistered, Taiwan Fisheries Research Institute Fishery Researcher 1 sta.FR1–PHI–14–95,950927,435–451m,14°41–42′N,123°24–21′E,adultmale905mmTL,northernLuzon,Philippines. Centrophorus tessellatus: MCZ 1031S (holotype), adult male875mmTL,SagamiBay,Japan,35°08′N,139°31′E, ca. 730 m, 09 Aug 1903.

Key to the Australian species of Centrophorus

1 Lateral trunk denticles with leaf-like flattenedcrowns on elevated narrow to broad pedicels extending above their bases; denticle crowns with strong medial and lateral cusps on their posterior ends ...................................................................... 2

Lateral trunk denticles with flat, sessile crownson their bases, without separate pedicels; denticle crowns sometimes with a posterior medial cusp but no lateral cusps ..................................................... 3

2 Free rear tips of pectoral fins broadly angularwithout an expanded lobe; lateral trunk denticles of adults leaf-like and on narrow pedicels

......................................................... C. squamosus (Indo–WestPacific,EasternAtlantic)

Free rear tips of pectoral fins expanded into ashort lobe; lateral trunk denticles of adults with a prominent angular cusp on a short, broad pedicel .. ................... C. acus (scatteredIndo–WestPacific)

3 Second dorsal fin relatively small, about halfheight of first dorsal fin; dorsal fins relativelywell spaced (interdorsal space >24% TL), second dorsal-spine origin behind pelvic-fin free reartip ................... C. moluccensis(Indo–WestPacific)

Second dorsal fin relatively large, about threequarters height of first dorsal fin; dorsal finsrelatively close together (interdorsal space <23% TL), second dorsal-spine origin usually above pelvic-fininnermargin ......................................... 4

4 Free rear tips of pectoral fins moderatelyelongate, usually not extending past first dorsalspine; lateral trunk denticles with somewhat elongated crowns and a prominent angular cusp in adults; large species, attaining >150 cm TL…..

................................ C. niaukang (= C. granulosus sensu Last & Stevens, 1994; Indo–West Pacific)

Free rear tips of pectoral fins greatly elongated,extending past first dorsal spine; lateral trunkdenticles with rounded or oval crowns and a low obtusely angular cusp; medium sized species, attaining ca. 110 cm TL........................................ 5

5 Dorsal fins with broad, dark greyish to blackishmargins (less distinct in adults but still obvious when fresh), without distinctive whitish markings; snout moderately long (depth at front of mouth less than 1.9 times preoral length; mouth width less than 1.2 in preoral length) ............................................... .............. C. zeehaani sp. nov. (southern Australia)

Dorsalfinswithdarkanteriorblotch(lessobviousin adults) and broad white posterior margin (much narrower in adults but still obvious when fresh); snout long (depth at front of mouth more than 2.0 times preoral length; mouth width more than 1.3 in preoral length) ...................................................... 6

6 Dorsalfinswidelyspaced(interdorsalspacemorethan 2.8 times dorsal–caudal space); relatively narrow head (width at front of mouth less than 10% of total length); mouth width more than 3.1 in head length ...................................................................... ........ C. westraliensis sp. nov. (Western Australia)

Dorsal fins close together (interdorsal space lessthan 2.6 times dorsal–caudal space); relatively broad head (width at front of mouth more than 10% of total length); mouth width less than 3.0 in head length ................................................ C. harrissoni

(Australia, New Zealand, New Caledonia)

20

ACKNOWLEDGEMENTS

ThisresearchprojectwassupportedbytheWealthfromOceans (WFO) Flagship. We thank the curators and curatorial staff for assistance with obtaining specimens and collection data: Alastair Graham (CSIRO), Di Bray (NMV), Mark McGrouther (AMS), Jeff Johnson (QM) and Ann Holmes and Karsten Hartel (MCZ). Special thanks go to Peter Last and John Stevens (CSIRO) for their invaluable contribution to the knowledge of Australia’s shark and ray fauna and their important preliminary taxonomicresearchonAustraliandogfishsharks.Thanksalso go to Ken Graham (NSW Department of Primary Industries) for providing images and specimens, Ross Daley (CSIRO) for collection of important type material and his ongoing research on Centrophorus sharks, Rory McAuley (Department of Fisheries, WA) for providing capture information and images of some Western Australian specimens, Louise Conboy and Thor Carter (CSIRO) for photographing of specimens, Tim Fountain (CSIRO) for preparing radiographs and obtaining meristic information, John Pogonoski (CSIRO) for checking meristic information and editorial comments, Dan Gledhill (CSIRO) for his technical assistance and Bob Ward and Bronwyn Holmes (CSIRO) for their extensive work on molecular barcoding of Australian sharks and rays. Themajorityof thespecimensof thenewspecieswerecollected by the FV Diana and FRV Southern Surveyor during surveys in Australian waters and we thank the crew and participants of those surveys. Dave Ebert would like to thank NOAA/NMFS for funding support of the National SharkResearchConsortiumandPacificSharkResearchCenter. L.J.V. Compagno would like to thank the National Research Foundation (South Africa), Iziko-South African Museum (Cape Town), Marine and Coastal Management (Cape Town), and the South African Institute of Aquatic Biodiversity (Grahamstown) for support including collection and curation of specimens. Indonesian material wascollectedaspartofa5-yearACIARfundedprojectandimportantcontributorstothisprojectincludeFahmi(Indonesian Institute of Sciences, LIPI), Dharmadi (Research Centre for Capture Fisheries, RCCF), Junaedi (Kedonganan fishmarket, Bali), processors and fishersat Tanjung Luar fish market (Lombok), Jenny Giles(Queensland Department of Primary Industries and Fisheries, QDPI&F), Ono Kurnaen Sumadhiharga (LIPI), Subhat Nurhakim (RCCF), Ian Potter (Murdoch University) and Steve Blaber (CSIRO).

REFERENCES

Anon. (1998) New sharks from New Caledonia. French Science and Technology 44: 10.

Baranes, A. (2003) Sharks from the Amirantes Islands, Seychelles, with a description of two new species of squaloids from the deep sea. Israel Journal of Zoology 49: 33–65.

Bleeker, P. (1860) Elfde bijdrage tot de kennis dervischfauna van Amboina. Acta Soc. Sci. Indo-Neerl. 8: 1–14.

Bloch, M.E. & Schneider, J.G. (1801) Systema ichthyo-logiae iconibus ex illustratum. Berlin, vol. 2, 584 pp.

Bocage, J. V. Barbosa du & Capello, F. de Brito (1864) Sur quelque espèces inédites de Squalidae de la tribu Acanthiana, Gray, qui fréquentent les côtes du Portugal. Proceedings of the Zoological Society of London 24: 260–263.

Bonnaterre, J.P. (1788) Tableau encyclopédique et méthodique des trois règnes de la nature. Ichthyologie. Paris, 215 pp.

Chu, Y.T., Meng, C.W. & Liu, J.X. (1981) Description of a new genus and a new species of Squalidae of China. Acta Zootaxon. Sinica 6: 100–103.

Compagno, L.J.V. (1984) FAO Species Catalogue. Vol. 4, Sharks of the World. An annotated and illustrated catalogue of shark species known to date. FAO Fisheries Synopsis No. 125. vol. 4, pt. 1 (noncarcharhinoids), pp. viii, 1–250.

Compagno, L.J.V. (2001) Sharks of the World: an annotated and illustrated catalogue of shark species known to date. Volume 2. Bullhead, mackerel and carpet sharks (Heterdontiformes, Lamniformes and Orectolobiformes). FAO, Rome, 269 pp.

Compagno, L.J.V., Dando, M. & Fowler, S. (2005) A Field Guide to Sharks of the World. Harper Collins Publishing Ltd., London, 368 pp.

Daley, R., Stevens, J.D., Last, P. & Yearsley, G. (2002) Field Guide to Australian Sharks and Rays. CSIRO Marine Research and Fisheries Research & Development Corporation, Australia, 84 pp.

Duffy, C.A.J. (2007) First record of Centrophorus harrissoni from New Zealand, with observations on squamation in Centrophoridae (Squaliformes). New Zealand Journal of Marine and Freshwater Research 41: 163–173.

Garman, S. (1906) New Plagiostomia. Bulletin of the Museum of Comparative Zoology 46: 203–208.

Garman, S. (1913) The Plagiostomia (sharks, skates, and rays). Memoirs of the Museum of Comparative Zoology, Harvard College 36: 515 pp.

Gomon, M.F., Glover, J.C.M. & Kuiter, R.H. (1994) The Fishes of Australia’s South Coast. State Print, Adelaide, 992 pp.

21Descriptions of new Australian chondrichthyans

Graham, K.J., Andrew, N.L. & Hodgson, K.E. (2001) Changes in relative abundance of sharks and rays on Australian South East Fishery trawl grounds after twenty years of fishing.Marine and Freshwater Research 52: 549–561.

Last, P.R. & Harris, J.G.K. (1981) New locality records and preliminary information on demersal fish faunalassemblages in Tasmanian waters. The Papers and Proceedings of the Royal Society of Tasmania 115: 189–209.

Last, P.R., White, W.T., Pogonoski, J.J., Gledhill, D.C., Yearsley, G.K. & Ward, R.D. (2007) Part 1 — Application of a rapid taxonomic approach to the genus Squalus, pp. 1–10. In: P.R. Last, W.T. White & J.J. Pogonoski (eds) Descriptions of new dogfishes of the genus Squalus (Squaloidea: Squalidae). CSIRO Marine and Atmospheric Research Paper 014, 130 pp.

Last, P.R. & Stevens, J.D. (1994) Sharks and Rays of Australia. CSIRO, Australia, 513 pp.

McCulloch,A.R.(1915)Reportonsomefishesobtainedby the F. I. S. “Endeavour” on the coasts of Queensland, New South Wales, Victoria, Tasmania, South and

South-Western Australia, Part III. Biological Results “Endeavour” 3: 97–170.

Müller, J. & Henle, F.G.J. (1837) Gattungen der HaifischeundRochennacheinervonihmmitHrn.Henleunternommenen gemeinschaftlichen Arbeit über die Naturgeschichte der Knorpelfische. Berichte Akademie der Wissenschaften zu Berlin 2: 111–118.

Muñoz-Chápuli, R. & Ramos, F. (1989) Review of the Centrophorus sharks (Elasmobranchii, Squalidae) of the eastern Atlantic. Cybium 13: 65–81.

Rafinesque,C.S.(1810)Caratteri di alcuni nuovi generi e nuove specie di animali e piante delta Sicilia. Palermo, 105 pp.

Teng, H.T. (1959) Studies on the elasmobranch fishesfrom Formosa. Pt. 6. A new species of deep-sea shark (Centrophorus niaukang) from Formosa. Report of the Laboratory of Fishery Biology, Taiwan, 9, 1–6.

Yearsley, G.K., Last, P.R. & Ward, R.D. (2001) Australian Seafood Handbook: an identification guide to domestic species. FRDC / CSIRO Marine Research, 469 pp.